Cárate Tandalla, D. (2016): Effects of moderate Nitrogen and Phosphorus addition on the species composition and dynamics of the tree seedlings community in tropical montane forests in southern Ecuador University of Goettingen, phd thesis

Abstract:

Anthropogenic activities have produced changes in natural ecosystems worldwide. In tropical regions in South America, industrialization of cities and forest clearance via burning are the main activities releasing pollutants into the atmosphere and inducing changes in nutrient deposition patterns and climate of primary forests.
Nitrogen (N) and phosphorus (P) are considered the main limiting nutrients of plant growth since their availability is vital for net primary productivity. Therefore, any change in N or P availability in soils would likely alter important mechanisms of forests dynamics such as growth and survival. Changes in soil pH (acidity), nutrient cycles and altered nutrient stocks affect N and P availability and affect various physiological processes of trees. Although low levels of nutrient deposition have been reported for montane forests in southern Ecuador (ca. 5 kg ha–1 for N, 0.49 kg ha–1 for P), even these levels are expected to lead to changes in forest structure and dynamics over the long term.
The responsiveness of forest to changes in resource availability varies with forest age and successional status, as well as life stage of the plant. Young plants (tree seedlings) should be more responsive to changes in nutrient availability than mature forest trees. Further, any demographic response is more likely to be visible in seedlings before mature trees because of the more rapid dynamics of seedlings. Therefore, I studied the regeneration dynamics of montane forest to understand which nutrient-related processes are involved in the growth and establishment of seedlings at both the individual and community levels.
The Ecuadorian NUtrient Manipulation EXperiment (NUMEX) has been designed to study the response of montane forest to moderate fertilization. The experiment has been set up over an elevation range across three main study sites (1000 m. a.s.l: Bombuscaro; 2000 m. a.s.l: San Francisco; 3000 m. a.s.l: Cajanuma) within the Podocarpus National Park and San Francisco Reserve. The factorial experiment consists of four blocks containing four experimental plots (N, P, NP and control) in every study site (16 plots per elevation). Fertilization has been done since 2008, adding moderate quantities of fertilizer (50 kg ha–1 y–1 of N and/or 10 kg ha–1 y–1 of P). Three different approaches were used to assess the seedling community and common species responses to fertilization. First, the seedling community was monitored in natural forest at 2000 m over three consecutive years (2011 – 2013). In 192 monitoring subplots (1m2 each) in San Francisco, all seedlings and saplings were mapped and tagged. Measurements of height, diameter, herbivory and leaf production were recorded for all individuals, and the number of recruited and dead seedlings was counted every year.
Second, allocation patterns and stoichiometry of seedlings of the six most common species were determined across the elevation gradient. Naturally occurring seedlings of the following species were harvested: Clarisia racemosa and Pouteria torta in Bombuscaro, Graffenrieda emarginata and Palicourea angustifolia in San Francisco and Grafferieda harlingii and Hedyosmum purpuracens in Cajanuma. Morphology (biomass allocation, herbivory and foliar areas such as SLA, LAR and LA) and foliar stoichiometry (nutrient contents and N:P ratios) were recorded and analyzed.
Third, a reciprocal transplantation experiment (STE) within the NUMEX experimental plots monitored seedlings of the most common species in Bombuscaro, (Pouteria torta), to assess specific responses in seedling performance over one year.
These three levels of analysis showed differential responses of the species community and common species to fertilization. Contrary to lowland forests, the community seedlings in this montane forest showed a moderate response. The density of seedling individuals decreased following nutrient addition, as a result of lower recruitment in treatments plots, but mortality was unaffected. Consequently, density-dependent mechanisms activated by additional N uptake did not show evidence of being a main driver to changes in species density.
The community of seedlings did not show significant growth in response to fertilization although seedlings were higher in the N treatment. However, N addition increased herbivory over all seedlings and plots, which could have masked other seedlings growth responses.
Common species seem to be well adapted to the relatively poor soils since these species were not favored by nutrient addition showing no change in growth or leaf traits. Over the long-term, common species might lose their dominance by being less competitive than faster growing species. However, it would need a longer period of monitoring under continued fertilization to produce visible shifts in community composition.
Species-specific responses are hard to determine in such species rich communities. Therefore, analyses of the six common species were conducted to complement the community-level study. Leaf morphology and foliar stoichiometry were assessed from harvested seedling from the experimental plots. Both N and P had effects on seedling traits, suggesting co-limitation of N and P in montane tree species in all elevations. However, foliar N:P ratios and the more frequently response to P addition indicated this nutrient might be more limiting than N along the gradient.
Responsiveness of the studied species varied between the six species. Stronger foliar P concentration compared with N in all species but Pouteria torta suggested higher P consumption in montane forest species, similar to the responses of several other tropical plant species after P addition. Increased herbivory was only evident in the opposite responses of G. emarginata (N and N+P addition) and P. angustifolia (P addition) at 2000 m suggesting that plant selection by herbivores is driven by resource quality. Most of the species had higher belowground biomass in root fractions following fertilization, except for G. harlingii at 3000 m, the only species that showed significant increase in aboveground biomass after nutrient addition.
Pouteria torta seedlings showed no significant changes in species performance after fertilization one year after establishment of the STE. Pouteria seems to be well-adapted to poor soils, since N and P addition did not alter foliar nutrient concentrations. Nevertheless, N and N+P addition significantly increased mortality and diameter growth rates. P addition resulted in higher leaf area loss and shifted carbon allocation to root growth. These responses indicated consequences in the competitive strength in the understory and recruitment success of Pouteria torta. However, the ambiguous response in some attributes (growth and herbivory) not related with mortality made it difficult to predict the future abundance of this species in long term.
In conclusion, complementary studies demonstrated that specific nutrient limitation for N or P in montane species seem not to be a rule in rich species ecosystems although nutrient addition did alter some pattern of growth and survival. Thus, nutrient fertilization might affect various mechanisms and dynamics of plant communities, the balance of which will only play out over long time scales.

Abstract:

Trait response-effects are critical to forecast community structure and biomass production in highly diverse tropical forests. Ecological theory and few observation studies indicate that trees with acquisitive functional traits would respond more strongly to higher resource availability than those with conservative traits. We assessed how long-term tree growth in experimental nutrient addition plots (N, P and N+P) varied as a function of morphological traits, tree size and species identity. We also evaluated how trait-based responses affected stand scale biomass production considering the community structure. We found that tree growth depended on interactions between functional traits and the type or combination of nutrients added. Common species with acquisitive functional traits responded more strongly to nutrient addition, mainly to N+P. Phosphorous enhanced the growth rates of species with acquisitive and conservative traits, had mostly positive effects on common species and neutral or negative effects in rare species. Moreover, trees receiving N+P grew faster irrespective of their initial size relative to control or to other treatment plots. Finally, species responses were highly idiosyncratic suggesting that community processes including competition and niche dimensionality may be altered under increased resource availability. We found no statistically significant effects of nutrient additions on aboveground biomass productivity because acquisitive species had a limited potential to increase their biomass, possibly due to their generally lower wood density. In contrast, P addition increased the growth rates of species characterized by more conservative resource strategies (with higher wood density) that were poorly represented in the plant community. We provide the first long-term experimental evidence that trait-based responses, community structure, and community processes modulate the effects of increased nutrient availability on biomass productivity in a tropical forest.

Abstract:

Although often overlooked in forest research, the canopy can play an important role in forest nutrient cycling. Since the canopy is spatially isolated from the forest floor, nutrient cycling in the
two areas may differ as terrestrial nutrients accumulate. We measured rates of free-living N2 fixation
along an elevation gradient (1,000, 2,000 and 3,000 m) of tropical montane canopy soils, compared
these to rates measured in the top 5 cm of forest floor soils (excluding fresh litter), and assessed the effects of elevated nutrient inputs to the forest floor. N2 fixation was measured using the acetylene reduction assay. Measurements occurred in the field, in the wet and dry seasons, using intact cores of soil. The forest floor had been fertilized biannually with moderate amounts of nitrogen (N) and phosphorus (P) for 4 years; treatments included control, N, P and N x P. N2 fixation rates exhibited little variation with Elevation but were higher in the dry season than the wet season. Fixation was inhibited in forest floor N plots compared to control and P plots, and stimulated in canopy P plots compared to control. At 2,000 m, the canopy contributed 12 % of measured canopy and forest floor N2 fixation (1.2 kg N ha-1 year-1).
Results suggest that N2 fixation is an active process in canopy soils, which is variable across seasons and
sensitive to changes in terrestrial nutrient availability. Long-term terrestrial accumulation of N and/or P has the potential to significantly change the dynamics of soil N cycling in these canopies.

Abstract:

Tropical ecosystems have an important role in global change scenarios, in part because they serve as a large terrestrial carbon pool. Carbon protection is mediated by soil aggregation processes, whereby biotic and abiotic factors influence the formation and stability of aggregates. Nutrient additions may affect soil structure indirectly by simultaneous shifts in biotic factors, mainly roots, and fungal hyphae, but also via impacts on abiotic soil properties. Here, we tested the hypothesis that soil aggregation will be affected by nutrient additions primarily via changes in arbuscular mycorrhizal fungal (AMF) hyphae and root length in a pristine tropical forest system. Therefore, the percentage of water-stable macroaggregates (> 250 ?m) (WSA) and the soil mean weight diameter (MWD) was analyzed, as well as nutrient contents, pH, root length, and AMF abundance. Phosphorus additions significantly increased the amount of WSA, which was consistent across two different sampling times. Despite a positive effect of phosphorus additions on extra-radical AMF biomass, no relationship between WSA and extra-radical AMF nor roots was revealed by regression analyses, contrary to the proposed hypothesis. These findings emphasize the importance of analyzing soil structure in understudied tropical systems, since it might be affected by increasing nutrient deposition expected in the future.

Abstract:

Nutrient deposition to tropical forests is increasing, which could affect soil fluxes of nitrous oxide (N2O), a powerful greenhouse gas. We assessed the effects of 35–56 months of moderate nitrogen (N) and phosphorus (P) additions on soil N2O fluxes and net soil N-cycling rates, and quantified the relative contributions of nitrification and denitrification to N2O fluxes. In 2008, a nutrient manipulation experiment was established along an elevation gradient (1000, 2000, and 3000 m) of montane forests in southern Ecuador. Treatments included control, N, P, and N+P addition (with additions of 50 kg N ha?1 yr?1 and 10 kg P ha?1 yr?1). Nitrous oxide fluxes were measured using static, vented chambers and N cycling was determined using the buried bag method. Measurements showed that denitrification was the main N2O source at all elevations, but that annual N2O emissions from control plots were low, and decreased along the elevation gradient (0.57 ± 0.26–0.05 ±0.04 kg N2O-N ha?1 yr?1). We attributed the low fluxes to our sites' conservative soil N cycling as well as gaseous N losses possibly being dominated by N2. Contrary to the first 21 months of the experiment, N addition did not affect N2O fluxes during the 35–56 month period, possibly due to low soil moisture contents during this time. With P addition, N2O fluxes and mineral N concentrations decreased during Months 35–56, presumably because plant P limitations were alleviated, increasing plant N uptake. Nitrogen plus phosphorus addition showed similar trends to N addition, but less pronounced given the counteracting effects of P addition. The combined results from this study (Months 1–21 and 35–56) showed that effects of N and P addition on soil N2O fluxes were not linear with time of exposure, highlighting the importance of long-term studies.

Abstract:

Ecosystems worldwide face increasing nutrient depositions mainly caused by anthropogenic processes. In particular, tropical ecosystems react sensitively to altering nutrient supply. The deposition of nutrients might influence the nutrient cycles, primarily of N and P in tropical montane rainforests. Increased nutrient supply leads to an enhanced biomass production and therefore other nutrients become limited for plants and microorganisms.
For this reason, the aim of this thesis is to study the response of phosphatase activity (PA) on moderate fertilization along an altitudinal gradient in a tropical montane rainforest in South Ecuador. The experiment was conducted on the NUMEX study sites including three different elevation levels 1000, 2000 and 3000m a.s.l. The different plots were treated with N, P, N+P to simulate increased nutrient depositions and one control plot. Further, organic layer and mineral soil was sampled and phosphomono- and phosphodiesterase activity (PMEA and PDEA) were determined.
The N fertilized plots showed only small effects compared to the control; presumably due to low amounts of added fertilizer. PA in the P addition plots showed reduced activity compared to the control with significant results of PMEA in the organic layer of the study sites on 2000 and 3000m a.s.l. The reason might be sufficient quantities of inorganic P which suppresses the production of phosphatases. Further, PA in N+P plots showed lower PA compared to the control than in the P addition plots. This effect could be caused through the dominating inhibitory effect of P in contrast to the stimulating effect of N on PA. Altitudinal differences were observed comparing the control plots at different elevation levels (1000, 2000 and 3000m a.s.l.). The results for the organic layer showed increasing PA along the altitudinal gradient with the lowest PA at 1000m a.s.l. The findings are contrary to the present literature which states that lowland tropical forests are characterized by high decomposition rates coming up with high PA. The findings cannot support this hypothesis; therefore more research is needed in the studied area in South Ecuador.

Abstract:

Tank bromeliads are common epiphytic plants throughout neotropical forests that store
signi?cant amounts of water in phytotelmata (tanks) formed by highly modi?ed leafs.
Methanogenic archaea in these tanks have recently been identi?ed as a signi?cant source of
atmospheric methane. We address the effects of environmental drivers (temperature, tank
water content, sodium phosphate [P], and urea [N] addition) on methane production in
anaerobically incubated bromeliad slurry and emissions from intact bromeliad tanks in
montane Ecuador. N addition ? 1 mg g 1 had a signi?cantly positive effect on headspace
methane concentrations in incubation jars while P addition did not affect methane
production at any dosage (? 1 mg g 1 ). Tank bromeliads (Tillandsia complanata) cultivated
in situ showed signi?cantly increased ef?uxes of methane in response to the addition of
26 mg N addition per tank but not to lower dosage of N or any dosage of P (? 5.2 mg plant 1 ).
There was no signi?cant interaction between N and P addition. The brevity of the
stimulatory effect of N addition on plant methane ef?uxes (1–2 days) points at N
competition by other microorganisms or bromeliads. Methane ef?ux from plants closely
followed within-day temperature ?uctuations over 24 h cycles, yet the dependency of
temperature was not exponential as typical for terrestrial wetlands but instead linear. In
simulated drought, methane emission from bromeliad tanks was maintained with minimum
amounts of water and regained after a short lag phase of approximately 24 h. Our results
suggest that methanogens in bromeliads are primarily limited by N and that direct effects of
global change (increasing temperature and seasonality, remote fertilization) on bromeliad
methane emissions are of moderate scale.

Abstract:

Nitrogen deposition to tropical forests is predicted to increase in future in many regions due to agricultural intensification. We conducted a seedling transplantation experiment in a tropical premontane forest in Ecuador with a locally abundant late-successional tree species (Pouteria torta, Sapotaceae) aimed at detecting species-specific responses to moderate N and P addition and to understand how increasing nutrient availability will affect regeneration. From locally collected seeds, 320 seedlings were produced and transplanted to the plots of the Ecuadorian Nutrient Manipulation Experiment (NUMEX) with three treatments (moderate N addition: 50 kg N ha?1 year?1, moderate P addition: 10 kg P ha?1 year?1 and combined N and P addition) and a control (80 plants per treatment). After 12 months, mortality, relative growth rate, leaf nutrient content and leaf herbivory rate were measured. N and NP addition significantly increased the mortality rate (70 vs. 54% in the control). However, N and P addition also increased the diameter growth rate of the surviving seedlings. N and P addition did not alter foliar nutrient concentrations and leaf N:P ratio, but N addition decreased the leaf C:N ratio and increased SLA. P addition (but not N addition) resulted in higher leaf area loss to herbivore consumption and also shifted carbon allocation to root growth. This fertilization experiment with a common rainforest tree species conducted in old-growth forest shows that already moderate doses of added N and P are affecting seedling performance which most likely will have consequences for the competitive strength in the understory and the recruitment success of P. torta. Simultaneous increases in growth, herbivory and mortality rates make it difficult to assess the species' overall performance and predict how a future increase in nutrient deposition will alter the abundance of this species in the Andean tropical montane forests.

Abstract:

In the past two decades, the tropical montane rain forests in south Ecuador experienced increasing deposition of reactive nitrogen mainly originating from Amazonian forest fires, while Saharan dust inputs episodically increased deposition of base metals. Increasing air temperature and unevenly distributed rainfall have allowed for longer dry spells in a perhumid ecosystem. This might have favored mineralization of dissolved organic matter (DOM) by microorganisms and increased nutrient release from the organic layer. Environmental change is expected to impact the functioning of this ecosystem belonging to the biodiversity hotspots of the Earth.
In 2007, we established a nutrient manipulation experiment (NUMEX) to understand the response of the ecosystem to moderately increased nutrient inputs. Since 2008, we have continuously applied 50 kg ha-1 a-1 of nitrogen (N), 10 kg ha-1 a-1 of phosphorus (P), 50 kg + 10 kg ha-1 a-1 of N and P and 10 kg ha-1 a-1 of calcium (Ca) in a randomized block design at 2000 m a.s.l. in a natural forest on the Amazonia-exposed slopes of the south Ecuadorian Andes.
Nitrogen concentrations in throughfall increased following N+P additions, while separate N amendments only increased nitrate concentrations. Total organic carbon (TOC) and dissolved organic nitrogen (DON) concentrations showed high seasonal variations in litter leachate and decreased significantly in the P and N+P treatments, but not in the N treatment. Thus, P availability plays a key role in the mineralization of DOM. TOC/DON ratios were narrower in throughfall than in litter leachate but their temporal course did not respond to nutrient amendments.
Our results revealed an initially fast, positive response of the C and N cycling to nutrient additions which declined with time. TOC and DON cycling only change if N and P supply are improved concurrently, while NO3-N leaching increases only if N is separately added. This indicates co-limitation of the microorganisms by N and P. The current increasing reactive N deposition will increase N export from the root zone, while it will only accelerate TOC and DON turnover if P availability is simultaneously increased. The Saharan dust-related Ca deposition has no impact on TOC and DON turnover.

Abstract:

Tropical montane forests of the Andes belong to the hotspots of biodiversity. But these nutrient poor ecosystems are simultaneously threatened by increased element inputs of nitrogen and phosphor in nutrient cycles. NUMEX-Projekt simulates the expected nutrient inputs in the for-ests and quantifies the changes of this ecosystem. In line of this study litter samples were collected by littertraps during a working period from February till May 2016. Leaf litter produc-tion, parameters of leaf morphology (leaf area and specific leaf area) and litter nutrients of nitrogen and phosphor were determined. Additionally, the parameters of herbivory (leaf mass loss and leaf area loss) were calculated by measuring holes area. The variation of these pa-rameters along an altitudinal gradient from 1.000 m.a.s.l. to 3.000 m.a.s.l. was investigated. Besides the variation after nitrogen and/or phosphor addition was explored. Furthermore, the extend of nutrients, lost through herbivory, was identified for the stand level. Terminatory the influence of two soil parameters (C/N ratio and Presin) was discussed.
There is a significant influence of altitude (p<0,05) on leaf morphology (leaf area: 1.000 m.a.s.l: 26,4±1,6 cm², 3.000 m.a.s.l: 7,4±1,0 cm², specific leaf area: 1.000 m.a.s.l: 99,7±4,1 cm²·g-1, 3.000 m.a.s.l: 56,1±6,6 cm²·g-1), leaf nutrients of nitrogen (1.000 m.a.s.l: 15,3±0,5 g·kg-1, 3.000 m.a.s.l: 6,6±0,7 g·kg-1) and phosphor (1.000 m.a.s.l: 0,54±0,03 g·kg-1, 3.000 m.a.s.l: 0,25±0,03 g·kg-1) and also on the parameters of herbivory: holes area (1.000 m.a.s.l: 1,8±0,2 cm², 3.000 m.a.s.l: 0,3±0,1 cm²) and leaf area loss (1.000 m.a.s.l: 6,8±0,4 %, 3.000 m.a.s.l: 3,6±0,6 %).
Nutrient addition did not lead to distinct results according to the variation of measured param-eters. NP fertilization had a positive effect on leaf area and P fertilization a positive effect on specific leaf area. Leaf nutrients showed various answers. For this parameter, NP fertilization causes significantly raised nitrogen and phosphor concentrations in litter in each site. Leaf litter production in 1.000 m.a.s.l site significantly decreased through NP fertilization. Referring to herbivory only a little number of significant effects were detected. Holes area increased on 2.000 m.a.s.l site by NP addition while leaf area loss in 1.000 m.a.s.l site – as well as leaf area production – decreased. On stand level, there was no variation of leaf area loss.
Soil parameters have a higher impact on measured parameters on stand level. Specific leaf area and leaf area loss correlate positively with soil C/N of upper mineral soil. Correlations of these parameters with plant available phosphor Presin is negative.
Nutrient losses on stand level decrease with altitude. There was a negative effect of NP addition on nitrogen loss in 1.000 m.a.s.l site and a positive effect of NP addition on phosphor loss in all sites.

Abstract:

Ecosystem services, provided from tropical forests, are indispensable for human beings. Coherencies in the system, their networks, drivers and various underlying pathways are not completely understood yet. Elucidation on directions of key nutrients and changes in organisms delivers the opportunity to get an overview about these relations.
Leaf functional traits are one important component to uncover those cascades and organizations. With their fast respond to environmental conditions, changes can be detected. Since, tropical regions suffer from increasing atmospheric inputs of nitrogen (N) and phosphorus (P), it is a need to investigate these progressions and predict future scenarios. The alarming fact, that these anthropogenic caused input have unknown consequences for the structure and functioning of tropical forests leads to a crucial study topic. Furthermore, questions should be answered if these increased inputs have a comparable influence on different elevations, based on identification the limiting nutrients.
To forecast these impacts the Nutrient Manipulation Experiment (NUMEX) has been conducted since 2008. It gained to show, that Neotropical montane forests respond rapidly to moderate nutrient additions of N (50 kg ha-1 yr-1) and P (10 kg ha-1 yr-1).
Within the present study, the eight most abundant tree species from the NUMEX sites were analyzed to their leaf functional traits from three elevations (altitude above sea level) in an Ecuadorian montane forest. The premontane forest in Bombuscaro (1,000 m) contains the species Clarisia racemosa and Pouteria torta. Further, in San Francisco (2,000 m), Alchornea lojaensis, Graffenrieda emarginata, Hieronyma fendleri, and Myrcia sp., were selected. In Cajanuma (3,000 m), the upper montane forest contains Hedyosmum purpurascens and Weinmannia loxensis.
The fertilization effects were calculated, using a mixed effect model, including study year and treatment as fixed effects and block and individual as random effects.
The results show, that foliar N decrease over the years, whereas, Foliar P conversely increases. Further, the relation of nitrogen and phosphorus (N:P) is decreasing during the study years. Upwards trends for ?15N are best replicated for the San Francisco site. With the leaf trait nutrient resorption efficiency (NuR) for N and P a decline over the study years is proceeding. In addition, leaf area (LA) and specific leaf area (SLA) tend to decrease. In conclusive processes over the study years might be the result of microclimatic events.
The sites differ in that San Francisco and Cajanuma respond more similar in some leaf functional traits, compared to them from Bombuscaro. It can be deduced that changes in the analyzed leaf traits over the last eight years between the sites and species are not significantly different. Summarizing, results represent the emphasis on the considerable influence of nutrient addition with notably changing foliar nutrient and show that the predicted nutrient deposition will probably change the ecosystem dynamics sustained.

Abstract:

The tropical montane forests of the E Andean cordillera in Ecuador receive episodic Sahara dust inputs particularly increasing Ca deposition. We added CaCl2 to isolate the effect of Ca deposition by Sahara dust to tropical montane forest from the simultaneously occurring pH effect. We examined components of the Ca cycle at four control plots and four plots with added Ca (2 × 5 kg ha–1 Ca annually as CaCl2) in a random arrangement. Between August 2007 and December 2009 (four applications of Ca), we determined Ca concentrations and fluxes in litter leachate, mineral soil solution (0.15 and 0.30 m depths), throughfall, and fine litterfall and Al concentrations and speciation in soil solutions. After 1 y of Ca addition, we assessed fine-root biomass, leaf area, and tree growth. Only < 3% of the applied Ca leached below the acid organic layer (pH 3.5–4.8). The added CaCl2 did not change electrical conductivity in the root zone after 2 y. In the second year of fertilization, Ca retention in the canopy of the Ca treatment tended to decrease relative to the control. After 2 y, 21% of the applied Ca was recycled to soil with throughfall and litterfall. One year after the first Ca addition, fine-root biomass had decreased significantly. Decreasing fine-root biomass might be attributed to a direct or an indirect beneficial effect of Ca on the soil decomposer community. Because of almost complete association of Al with dissolved organic matter and high free Ca2+ : Al3+ activity ratios in solution of all plots, Al toxicity was unlikely. We conclude that the added Ca was retained in the system and had beneficial effects on some plants.

Abstract:

Large areas in the tropics receive elevated atmospheric nutrient inputs. Presently, little is known on how nitrogen (N) cycling in tropical montane forest soils will respond to such increased nutrient inputs. We assessed how gross rates of mineral N production (N mineralization and nitrification) and microbial N retention (NH4+ and NO3- immobilization and dissimilatory NO3- reduction to NH4+ [DNRA]) change with elevated N and phosphorus (P) inputs in montane forest soils at 1000-, 2000-, and 3000-m elevations in
south Ecuador. At each elevation, four replicate plots (20320 m each) of control, N (added at 50 kg N ha-1yr-1), P (added at 10 kg P ha-1 yr-1), and combined N x P additions have been established since 2008. We measured gross N cycling rates in 2010 and 2011, using 15N pool dilution techniques with in situ incubation of intact soil cores taken from the top 5 cm of soil. In control plots, gross soil-N cycling rates decreased with increase in elevation, and microbial N retention was tightly coupled with mineral N production. At 1000 m and 2000 m, four-year N and combined N þ P additions increased gross mineral N production but decreased NH4+ and NO3- immobilization and DNRA compared to the control. At 3000 m, four-year N and combined N x P additions increased gross N mineralization rates and decreased DNRA
compared to the control; although NH4+ and NO3- immobilization in the N and NxP plots were not different from the control, these were lower than their respective mineral N production. At all elevations, decreased microbial N retention was accompanied by decreased microbial biomass C and C:N ratio. P addition did not affect any of the soil-N cycling processes. Our results signified that four years of N addition, at a rate expected to occur at these sites, uncoupled the soil-N cycling processes, as indicated by decreased microbial N retention. This fast response of soil-N cycling processes across elevations implies that greater
attention should be paid to the biological implications on montane forests of such uncoupled soil-N cycling.

Abstract:

Atmospheric nutrient deposition and climate change are expected to endanger the diversity of tropical forest ecosystems. Nitrogen (N) deposition might influence nutrient fluxes beyond the N cycle by a concomitant increased demand for other nutritional elements such as phosphorus (P). Organisms might respond to the increased P demand by enhanced activity of enzymes involved in releasing inorganic P from organic matter (OM). Our aims were to assess the effect of i) climate shifts (approximated by an altitudinal gradient), and ii) nutrient addition (N, P, N+P) on phosphatase activity (PA) in organic layer and mineral soil of a tropical montane rainforest in Southern Ecuador. A nutrient manipulation experiment (NUMEX) was set up along an altitudinal gradient (1000, 2000, and 3000 m a.s.l.). We determined PA and inorganic and total P concentrations. PA at 1000 m was significantly lower (mean ± standard error: 48 ± 20 µmol p-NP g-1 dm h-1) as compared to 2000 m and 3000 m (119 ± 11 and 137 ± 19, respectively). One explanation might be that very rapid decomposition of OM at 1000 m results in very thin organic layers reducing the stabilization of enzymes and thus, resulting in leaching loss of enzymes under the humid tropical climate. We found no effect of N addition on PA neither in the organic layer nor in mineral soil, probably because of the low nutrient addition rates that showed ambiguous results so far on productivity measures as a proxy for P demand. In the organic layers of P and N+P treatments, we found decreased PA and increased concentrations of inorganic P. This indicates that the surplus of inorganic P reduced the biosynthesis of phosphatase enzymes. PA in megadiverse montane rainforests is likely to be unaffected by increased atmospheric N deposition but reduced upon atmospheric P deposition.

Abstract:

Global changes in nutrient deposition rates are likely to have profound effects on plant communities, particularly in the nutrient-limited systems of the tropics. We studied the effects of increased nutrient availability on the seedlings of six tree species in montane forests of southern Ecuador in situ. After five years of continued N, P, or N+P addition, naturally grown seedlings of each of the two most common
species at each elevation (1000, 2000, and 3000 m asl) were harvested for analyses of leaf morphology, nutrient content, herbivory, and tissue biomass allocation. Most species showed increased foliar N and P concentrations after addition of each respective element. Leaf tissue N:P ratios of >20 in the control plants of all species suggest that P is more growth-limiting in these forests than N. Leaf
morphological responses to nutrient addition were species and nutrient specific, with some species (Hedyosmum purparescens, Graffenrieda emarginata) exhibiting increased specific leaf area (SLA), and others (Graffenrieda harlingii) increased leaf area ratios (LAR). Pouteria torta (1000 m) had lower SLA and LAR after P addition. Increased herbivory was only evident in G. emarginata (after N and N+P addition).
Only the species from 3000 m asl modified biomass allocation after nutrient addition. In general, N and N+P addition more strongly affected the species studied at the upper elevations, whereas P addition had a similar range of effects on the species at all elevations. We conclude that the responses of the studied tropical montane forest tree seedlings to chronic N and P addition are highly species-specific and that successful adaptation to increased nutrient availability will depend on species-specific morphological and physiological plasticity.

Abstract:

Studies in temperate systems provide evidence that the abundance of arbuscular mycorrhizal fungal (AMF) depends on soil nutrient availability, which is mainly explained in the context of resource stoichiometry and differential plant biomass allocation. We applied this concept to an understudied ecosystem – tropical montane forest – analyzing root and AMF abundance along an elevational gradient with decreasing nutrient availability, combined with responses to nitrogen (N) versus phosphorus (P) additions. At three sites from 1000 to 3000 m above sea-level we analyzed fine root length, AMF root colonization as well as extraradical AMF biomass (neutral lipid fatty acid 16:1?5, hyphal length and spore counts) in a nutrient manipulation experiment. We found a significant increase in root length as well as intra- and extraradical AMF abundance with elevation. Overall, P additions significantly increased, whereas N additions decreased AMF abundance, with differential though nonsystematic changes along the elevational gradient. Strongest effects were clearly observed at the intermediate site. These findings suggest a general dependency of roots and AMF on nutrient availability, though responses to N and P additions differed from previous studies in temperate systems. In the context of future nutrient depositions, results suggest diverging responses of AMF abundance depending on site characteristics.